Genetic variability in the cosmopolitan deep-water ophiuran Ophiomusium lymani

A population of the cosmopolitan deep-sea ophiuran Ophiomusium lymani was studied by gel electrophoresis, and proved to be highly variable genetically; about 53% of the 15 loci studied are polymorphic, and the average individual is heterozygous at about 17% of the loci. This is approximately the same genetic variability displayed by other species, belonging to other phyla or classes, from the same deep-sea trawl. A similarly high level of genetic variation occurs in deep-sea organisms in general, and in a shallow-water tropical species. Both the deep-sea and the tropics are trophically stable environments. On the other hand, low genetic variabilities have been found in marine species from trophically unstable environments. These data suggest that any phylogenetic effects on genetic variability are secondary, and that the trophic regime may be of major importance in determining genetic strategies of adaptation.This study is no. 5 of a series of reports on the results of Expedition Quagmire.     

Comparative study of sub-cuticular bacteria in brittlestars (Echinodermata: Ophiuroidea)

Arm tissues from species of brittlestars from the British Isles and North America were examined by light and electron microscopy for the presence of sub-cuticular bacteria (SCB) which were recorded in 17 of the 19 species studied. Three major groups of SCB could be distinguished on the basis of their morphology. The bacterial morphology was not correlated with the host ecology but did appear to be related to the phylogeny of the hosts. Ophiuroids of the same genera had morphologically similar SCB and this was also generally true of species from the same family. This indicates the possibility of co-evolution between the SCB and their hosts. Estimates of SCB load were made for 9 of the species by direct counting and by quantifying the amount of bacterial endotoxin present in the host tissues. SCB numbers varied interspecifically from 3.7×10⁸ to 4.6×10⁹ SCB per gram of ash-free brittlestar arm tissue. Endotoxin values ranged from 5.97 to 285 g g^–1 ash-free arm tissue. Calculations suggest that SCB form at least 0.1 to 1% of the total biomass of the arm tissues. There was considerable intraspecific variation in SCB load. The possible role and significance of SCB is discussed.     

Reproduction in Amphiura filiformis (Echinodermata: Ophiuroidea): Seasonality in gonad development

The burrowing ophiuroid Amphiura filiformis (O. F. Müller) (Echinodermata: Ophiuroidea) colonises an extensive area and is a numerically dominant member of the macrofauna in Galway Bay on the west coast of Ireland. There, it has a discrete, relatively short annual breeding period, from June to September, with peak activity in the month of August. The breeding period approximates in timing those of other European populations. The oocytes of a particular year are laid down in the autumn of the previous year; they begin to develop in spring, with the period of fastest growth in May/June. Gonad growth corresponds quite closely to the annual rise in water temperature, with spawning taking place during the warmest months of the year. The species is polytelic, and it is suggested that the long-lived Galway Bay individuals may breed every year for several years. The relationship between spawning and population dynamics is discussed in the light of current, available literature concerning A. filiformis from other parts of Europe. The role of so-called ultimate factors in the process of ophiuroid maturation and spawning is discussed.     

Large-scale estimate of somatic growth in Amphiura filiformis (Echinodermata: Ophiuroidea)

Quantitative data on size structure of the ophiuroid Amphiura filiformis (O.F. Müller) from 35 benthic stations in the Kattegat sampled twice with a 143 d interval, June and October 1991, were used to estimate somatic growth. The material was objectively divided into cohorts, and cohorts, from the two occasions were paired to give estimates of growth. The growth constant, K, in the Von Bertalanffy equation, was estimated from a Ford-Walford plot to 0.54 yr^-1. Results were in agreement with previous estimates from a few single sites, and suggest that the main part of dise growth occurs within the first 5 to 7 yr of living. Size specific growth in oral width was density independent despite high densities of A. filiformis (>3000 ind m^-2) and high total benthic biomass (up to 1000 g wet wt m^-2) in some areas. Growth was uncorrelated with plant pigment concentrations in the sediment and showed weak positive correlation with sediment carbon and nitrogen, as well as water depth. Growth was higher in fine sediments. This is the first attempt to estimate growth in this important species over a large area and to relate growth in the field to environmental factors.     

Post-metamorphic growth of the arms in Ophiophragmus filograneus (Echinodermata: Ophiuroidea) from Tampa Bay,Florida (USA)

Juveniles of the burrowing amphiurid Ophiophragmus filograneus from Tampa Bay, Florida (USA) exhibit a growth pattern unreported in the ophiuroid literature. Two nonadjacent arms grow at a greater rate than the other three arms. This phenomenon might be a developmental adaptation to avoid salinity and temperature fluctuations in surface waters of the shallow-water environment which O. filograneus inhabits. It is proposed that concentration of growth into 2 of the 5 arms permits earlier descent of the disc into the substratum with continued ability to feed on the surface with the arm tips. Larger specimens show a gradual equalization of arm lengths. Changes in growth rates and cropping of the longer arms by breakage and predation might account for such eventual equalization.     

Reproductive biology of a deep-sea brittle star Amphiura carchara (Echinodermata: Ophiuroidea)

 New information on the brooding reproduction of Amphiura carchara, and previous studies of several deep-sea congeners, contradict the prevailing notion that direct development is exceptional among deep-sea echinoderms. Over 500 specimens of A. carchara from 2,850 to 4,100-m depths off the coast of California, USA, were studied. The species was found to be gonochoric, although most brooding species of ophiuroids are hermaphroditic. Females each brooded up to 72 embryos at a time, with up to 10 in a single bursa. The embryos of individual adults were often at different stages of development, but those brooded in a single bursa tended to be at the same stage. Thus, the species has characteristics that breach the distinctions between sequential and simultaneous brooding exhibited by shallow-water ophiuroids. The embryos developed from yolky eggs that are large for an ophiuroid, with the mean largest oocytes 0.45 mm, and ranging to 1.28 mm in diameter. Almost all late-stage embryos were positioned with their mouth and arms pressed against the wall of the bursa, possibly to facilitate the uptake of nutrients from the parent. Thus, A. carchara may be matrotrophic. The largest embryos examined had a rudimentary disk skeleton, and arms with four joints and a terminal plate. Emerging juveniles probably differ in disk diameter. The species appears to brood year round, although differences in gonad size, the incidence of brooding, and the relative numbers of early developmental stages in summer and winter samples indicated that there are seasonal trends in reproduction. Received: 14 February 2000 / Accepted: 3 July 2000     

Reproduction and population dynamics of the fissiparous brittle star Ophiactis savignyi (Echinodermata: Ophiuroidea)

Reproductive cycle, asexual reproduction, and population dynamics of the fissiparous brittle star, Ophiactis savignyi, which inhabits the exhalant passages of the sponge Haliclona sp. were examined monthly from February 1991 to January 1992 at Wanlitung, southern Taiwan (22°N; 120°E). Mature gametes were found from March to December, but release was mainly in May and June. Sexual recruits were found from May to December, with the highest frequency (14.1% of population) in June. Sexual recruits composed 2.4% of the 1-yr sample. Sex ratio of male to female was 24:1. Fission occurred throughout the year, although the frequency of recently split individuals was lower from January to June (6 to 31%), and higher from July to December (42 to 52%). The occurrence of fission was highest after spawning. Regenerating individuals composed 48.2% of the 1-yr sample. Population density fluctuated greatly during summer due to recruitment by fission and mortality or dispersal due to the stressful environmental conditions. Both sexual and asexual reproduction of O. savignyi were successful at this site.     

The early embryology of the basket star Gorgonocephalus caryi (Echinodermata,Ophiuroidea)

The changes in morphology of Gorgonocephalus caryi embryos over the 8 1/2 day period of their survival are described. Fertilized eggs are 220 in diameter. Division is equal and holoblastic, and the blastula stage is reached within 24 h. Gastrulation begins at about 40 h. At 68 h, the anterior end of the embryo begins to elongate. The embryos are pear-shaped at 72 h. At about 84 h the anterior end begins to bend towards one side. By 98 h the entire embryo has flattened perpendicular to the anterior-posterior axis, and the embryo is a flattened, trilobed structure. At 4 1/2 days the anterior tip twists around to lie with its axis in a plane perpendicular to the other 2 lobes. They, meanwhile, shift in position so that the resulting embryo resembles an umbrella which is flat on top and has a curved handle. Meanwhile, the hydrocoel forms as an out-pouching from the archenteron in the anterior lobe. The general coelom develops by schizocoely. Between 5 and 5 1/2 days, the anterior lobe coelesces with the flattened area as it becomes pentagonal. By 5 1/2 days, many embryos are starshaped. At 6 1/2 days the mouth begins to form. By 8 days the embryos were dying, and none lived beyond 10 1/2 days. At no time did cilia or other locomotory structures develop. The development of G. caryi does not resemble that of any other ophiuroid studied. There is not trace of a pluteus, and it is postulated that the pluteus evolved after the Phrynophiurida and the Ophiurida were differentiated.     

Genetic variation in the neritic squid Loligo forbesi (Myopsida: Loliginidae) in the northeast Atlantic Ocean

Horizontal starch gel electrophoresis was employed to investigate levels of genetic differentiation between 13 samples of the neritic squid species Loligo forbesi Steenstrup obtained from throughout the majority of its known geographical range. Six enzyme loci identified in a preliminary study as being polymorphic were screened for variation between samples. No significant differences in allele distribution were detected between any of the samples obtained from the Faroe Bank in the north to Lisbon in the south, suggesting that squid throughout this range in the vicinity of the continental shelf are able to maintain panmixia, and effectively belong to a single population sharing a common gene pool. No clinal variation in allele distribution was detected throughout this range, a result which complements the findings of a detailed morphological companion study of the same individuals. Comparison of this homogenous European continental shelf population with squid from the Azores revealed highly significant (P<0.01) differences in allele distribution at five of the six polymorphic enzyme loci studied. A genetic identity value (I) equivalent to 0.93 over 33 loci was obtained. Analysis of F-statistics suggested migration rates between sites to be as low as one individual per five generations, a rate deemed insufficient under most models to prevent divergence by random genetic drift. The large distance and oceanic depths separating the Azores from continental Europe seem to present an effective barrier to gene flow to L. forbesi, a squid belonging to a family considered to be confined in distribution to relatively shallow, near coastal waters. The two populations of squid in the Azores and along the European continental shelf currently both ascribed to L. forbesi should therefore probably best be regarded as relative subspecies.     

Spawning behaviors of northeast Pacific sea cucumbers (Holothuroidea: Echinodermata)

Spawning behaviors observed from May 1981 through August 1983 in the laboratory and field are described and compared for twelve species of northeast Pacific holothuroids collected from waters of the San Juan Archipelago, Washington. For many species, elevation of the anterior end, stretching of the introvert, tentacle waving, and swaying are behaviors commonly associated with gamete release. Less common activities include waving of introvert podia, waves of body wall contractions, errant males, aggregations, and pseudocopulation. Positively buoyant eggs of four species [Psolus chitonoides Clark, Cucumaria miniata (Brandt), C. piperata (Stimpson), and C. fallax (Ludwig)] were packaged in pellets, ropes, or strands and released passively; a female Molpadia intermedia (Ludwig) discharged negatively buoyant eggs in a single explosive burst. Two externally brooding sea cucumbers [C. lubrica (Clark) and C. pseudocurata Deichmann] produced strands of dilution-resistant sperm that were carried by currents to nearby females. Three species commonly sympatric [P. chitonoides, C. miniata, and Eupentacta quinquesemita (Selenka)] in high-current rocky habitats have overlapping spring breeding seasons, but spawn at different hours of the day.Please address all requests for reprints to Dr. McEuen at the Harbor Branch Oceanographic Institution     

Effect of arm regeneration on energy storage and gonad production in Ophiocoma echinata (Echinodermata: Ophiuroidea)

Two laboratory experiments were conducted on Ophiocoma echinata (Lamarck, 1816) collected from Missouri Key, Florida, USA, to ascertain the effects of arm regeneration on energy storage and gonad production. In each experiment individuals were divided into non-regenerating (all arms intact) and regenerating (three arms autotomized) groups and maintained together in a re-circulating seawater system for 2 mo. In Experiment 1, individuals were fed once a week, and in Experiment 2 once every 2 wk. The amount of food energy required for maintenance was estimated to be 0.172 kJ d⁻¹. The lower feeding level (Experiment 2) was below maintenance ration and both groups lost material relative to an initial group. At the higher feeding level (Experiment 1), regeneration of three arms resulted in less storage of organic material, mainly lipid in the stomach, and less gonad production. Thus, regeneration may reduce energy reserves and reproductive output. Received: 23 April 1998 / Accepted: 11 January 1999     

Effects of salinity and calcium concentration on arm regeneration byOphiothrix angulata (Echinodermata: Ophiuroidea)

At 33 salinity a tissue stump formed 2 to 3 d after autotomy and developing ossicles were present by the fourth day inOphiothrix angulata (Say). Regeneration proceeded rapidly from the sixth day until the thirteenth day, when the rate decreased greatly. The length of the regenerated arm and the number of ossicles formed did not vary over a salinity range of 28 to 38 S, but were significantly less at 23 S. The number of ossicles regenerated increased linearly (y=1.9 x-7.7;r=0.9089) with the calcium concentrations ranging from 3.8 to 9.5 mM. No ossicle formation occurred at 3.8 mM calcium concentration. Rate of net uptake of calcium-45 into the ossicles of intact individuals in salinities of 28 and 33 was significantly greater than that in 23 and 38 S. However, net uptake rate of calcium into the soft tissues of the arms was significantly higher at 18S than at the lower two salinities.     

Tolerance to low oxygen and sulfide in Amphiura filiformis and Ophiura albida (Echinodermata: Ophiuroidea)

The brittle stars Amphiura filiformis (Müller) and Ophiura albida (Forbes) were exposed to different oxygen saturations (100, 10, 5, 3, and <1% oxygen saturation) and to physiological anoxia (<1% oxygen saturation) at different total sulfide concentrations (0, 2, 20, 200 μM). The mortality was followed during experiments and the median survival time (LT₅₀) was determined. The infaunal A. filiformis had a significantly higher tolerance to both hypoxia and sulfide than did the epibenthic O. albida. After exposure to 10% oxygen saturation for a month, only 2.0% A. filiformis and 0% O. albida were dead. In oxygen saturations <1% A. filiformis and O. albida had a LT₅₀ of 7.5 and 2.5 d, respectively. The presence of even very small concentrations of sulfide decreased the survival significantly. Sulfide is shown to be the key factor for the survival of the two species. Received: 11 October 1996 / Accepted: 12 November 1996     

Effects of disc and arm loss on regeneration byMicrophiopholis gracillima (Echinodermata: Ophiuroidea) in nutrient-free seawater

Clibanarius longitarsus (De Haan), a hermit crab, feeds by means of both micro- and macrofeeding. Microfeeding is of two types: filter feeding and deposit feeding. In macrofeeding, the animal depends mainly on green algae, barnacles and some worms, etc. In filter feeding, the hermit crab uses its paired antennules for trapping microorganisms drawn into the water current created by the second and third maxillipeds; the antennae do not play any role in filter feeding. In deposit feeding, the hermit crab uses its chelae as well as pereiopods for toughing and obtaining detritus food material. In macrofeeding, the organism employs sharp chelae to collect small pieces of muscle from barnacles and then transfer them to the inner mouth parts. In algal feeding, the chelae as well as maxillipeds help in collecting and moving food material towards the mouth. In this investigation functional organisation ofC. longitarsus mouth parts was studied in detail with reference to the various feeding mechanisms, in specimens collected from the intertidal region of Bheemunipatnam, Andra Pradesh, India, in 1988 and 1989.     

Population dynamics of Amphiura chiajei (Echinodermata: Ophiuroidea) in Killary Harbour,on the west coast of Ireland

The infaunal ophiuroid Amphiura chiajei Forbes is a dominant member of the A. chiajei community in Killary Harbour, a fjord-like inlet on the west coast of Ireland. High density populations (700 individuals/ m²) occur in sediments with a silt/clay content of 80 to 90% and organic carbon levels of 5 to 7%. A study of the population dynamics of this species was carried out from November 1985 to October 1988. At the outset, the population consisted of approximately equal numbers of adults and juveniles. By October 1988, the percentage of adults had increased to 95%, with little or no change in the mean annual dersities. However, there was some degree of variability between the mean monthly densities. Such variability may have been due to very low bottom temperatures, resulting in mortality among the older members of the population. Observations suggest that in Killary Harbour the species may attain an age of 10 yr, with an early annual growth rate of 0.5 mm (oral width). Annual recruitment was variable and low, due presumably to intraspecific competition with the adults; recruitment success may be very closely linked to adult mortality. The Killary Harbour findings are considered in the light of what is currently known of similar amphiurid populations in different geographical locations.     

Growth,production and ecological significance of Ophiura albida and O. ophiura (Echinodermata: Ophiuroidea) in the German Bight

Growth and production of the shallow-water ophiuroids Ophiura albida and O. ophiura were investigated at two stations in the German Bight from 1988 to 1991. Growth rings visible on the vertebral ossicles of the ophiuroid arms were interpreted as annual age markers. A correction for overgrown first rings allows for more exact estimations of growth and age. In both species growth could be described by Von Bertalanffy growth functions with the asymptotic disc diameter D =10.1 mm, K=0.229 and t _o=-0.192 in O. albida and D =27.7 mm, K=0.084 and t _o=0.042 in O. ophiura. Somatic production was calculated from mass specific growth rates. Annual production:biomass (P:B) ratios were estimated at 0.32 for O. albida and 0.43 for O. ophiura.AWI Publication Number: 618     

Field survey of the occurrence and significance of regeneration in Amphiura filiformis (Echinodermata: Ophiuroidea) from Galway Bay,west coast of Ireland

The Boreo-Mediterranean amphiurid Amphiura filiformis (O. F. Müller) occurs in high densities within Galway Bay. This burrowing species consistently shows signs of recent regeneration (of both arms and disc “cap”) at the above locality. Specimens were collected by SCUBA in March, April and May 1980, May 1981, and April 1982. Biomass assays have revealed that individuals with up to 50% (mean ca. 25%) of their entire body weight consisting of regenerated tissue are not uncommon. The occurrence of arm breakage and regrowth was found to be more frequent in the proximal (basal) and mid-arm regions than in the exposed arm tip portions. Roughly 3% of the population were found to be in the process of regenerating the disc “cap” or covering. Preliminary results from gut analyses of potential predators at the sampling station indicate that young flatfish (plaice and dab) actively “crop” the arms of A. filiformis. Records from north western Europe of this species occurring in the diets of fish and invertebrates are considered. In this long-lived species, the continual necessity to regenerate is deemed to be a major drain on bodily resources.     

Field survey of the occurrence and significance of regeneration in Amphiura chiajei (Echinodermata: Ophiuroidea) from Killary Habrour,west coast of Ireland

The boreo-Mediterranean amphiurid Amphiura chiajei Forbes occurs in high numbers (700 individuals/m²) in Killary Harbour, a fjordic inlet on the west coast of Ireland. 99.1% of the adult individuals show signs of arm regeneration, while 0.5% show regeneration of the disc. A study of the phenomenon commenced in November 1987 and continued on a seasonal basis until July 1989. Specimens were collected by SCUBA. The overall mean length of regenerated arm tissue over the 21 mo sampling period was 33.5±6.5 mm. Regeneration points were most common in the proximal and in the distal one-third of each arm. An overall mean of 4.21±0.3 arms per individual showed evidence of regeneration, while biomass assays revealed that regenerated tissue accounted for up to 57.9% of the total body weight. Anthropogenic activity (e.g. bottom trawling) may contribute to regeneration, although sub-lethal predation appears to be the main causitive factor of regeneration. Gut analyses of potential predators at the sampling station revealed that fish species (especially the Pleuronectidae) are particularly implicated. This predation seems to vary seasonally, being greatest in the summer months. Since the larger individuals showed the greatest evidence of arm regeneration, it is assumed that such sub-lethal damage is a persistent experience. Whatever the causality, the population of A. chiajei in Killary Harbour is deemed to be highly resilient.